The Subgeneric Classification of Species of Culicoides - thoughts and a warning

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1 Last revised: May 16, 2016 The Subgeneric Classification of Species of Culicoides - thoughts and a warning The present classification of species of Culicoides is, in my opinion, in terrible condition, such that for many years I simply listed all the species alphabetically in my world catalog (published in 1997 and regularly updated here on line). However, I hope that the following will promote further study, as we start to recognize monophyletic lineages. The arrangement below of species of the genus Culicoides is almost entirely phenetic (based on overall similarity). Amazingly, and in spite of the tremendous medical, veterinary and economic importance of so many species in the genus, there has been very little cladistic interpretation of the genus or of any group within the genus, based on morphology (i.e. interpretations of derived character states based on outgroup comparisons). Repeatedly, authors have classified species according to their similarity to one or another taxon, often noting that the antennal sensilla pattern is the same as one group, the wing another group and the male genitalia to yet another. It is clear that basing a classification on such similarities leads to phylogenetic chaos, and hence to continued instability in all future classifications. As such, there is a tremendous opportunity for future work to consider what might be derived within the genus (such as the spermathecal shape in Monoculicoides, the elongate spicules on the operculum of the pupae of species of Avaritia, etc.). A good approach by an ambitious researcher would be to take the type species of every described genus, the characteristic species of each species group, and a goodly smattering of other species, compare these carefully in alcohol, cleared in glycerine, and then on slides (without crushing various parts), obtain the larvae and pupae of as many representative taxa as possible, determine derived character states (based on outgroup comparisons) and combine the results to synthesize a phylogeny which would then provide the basis for further interpretation of species. It would be a gift to our science! At present it is uncertain whether most of the subgenera or species groups are monophyletic, although I believe some of them to be so, based on unpublished synapomorphies. Meiswinkel and Dyce (1989b) provided a rare and welcome overview of some character states of phylogenetic importance. Szadziewski et al. (2016) provided a synapomorphy for C. (Oecacta). Borkent (2014) and Shults et al. (2016) discuss synapomorphies of the genus as a whole. Aside from the species placed to subgenus, there are numbers placed in separate species groups of uncertain affiliation and a further list of miscellaneous species which no one knows what to do with. Aside from the theoretical problems noted above, this is also a reflection of various scientists working on their local fauna and constructing classifications as best they can on their local and inherently limited knowledge. There is a great need to study Culicoides on a worldwide basis, using

2 exemplars of the various groups from each region. A number of recent papers have used genomic methods to interpret phylogenetic relationships between select species of Culicoides (e.g. Ander et al., 2013;; Augot et al., 2013;; Bakhoum et al., 2013;; Lassen et al., 2012;; Linton et al., 2002;; Morag et al., 2012;; Schwenkenbecher et al., 2008;; Tay et al., 2016). Most use COI (or a couple of genes) as a measure but it is highly doubtful as to whether these genes (and especially COI on its own) can be used to interpret phylogenetic relationships. For example, low percentages of chironomid and simuliid genera and subgroups are recovered using COI barcodes (Ekrem et al. 2007, 2010, Rivera and Currie 2009). Further to this, readers are reminded that members of the genus Culicoides are known from the mid Cretaceous and there are 17 species known from all of the Cretaceous. The genus is clearly ancient and was certainly diverse even before the Cenozoic (see list of fossil species toward the end of this document). Further to this, very few barcoding papers indicate where their voucher specimens are housed indeed, many do not even keep vouchers, preventing future studies from being able to check published identifications. As such many of these will become throw-away studies because of uncertain species identification. This poor practice should be rectified immediately so that future researchers can reexamine those vouchers and interpret discrepancies. The immatures of the genus are poorly understood, in spite of the pioneering work on the larvae of a number of the Nearctic species by Murphree and Mullen (1991) and a number of smaller papers describing various larvae and pupae of select species. The literature describing the immatures (eggs, larvae, pupae) was listed in a recent publication by Borkent (2014, Table 2). This publication also provides the first generic keys to pupae and the first means of recognizing the pupae of Culicoides as such. In addition, pupal morphology provides important information for interpreting phylogenetic relationships. Currently, the larvae of 13% and the pupae of 17% of described species of Culicoides are known (but many poorly described). As a consequence, we don't know the habitat (or the features of the immatures) of the vast majority of named Culicoides. The following arrangement of species does NOT follow the most recent literature (which a good catalog should otherwise do), reflecting the chaos in our present classification. In instances of recent differences in opinion I have generally followed the opinions of Wirth who studied the genus worldwide. For example, the placement by Yu et al. (2006) of C. dendrophilus (type species of C. (Amossovia)), C. segnis (type species of C. (Wirthomyia)) and C. fascipennis (type species of C. (Silvaticulicoides)) in the subgenus C. (Oecacta) actually results in all the aforementioned subgenera becoming junior synonyms of the subgenus C. (Oecacta);; these subgeneric placements (without action on the subgeneric synonymy by Yu et al., 2006) have not been followed. In relation to this, Szadziewski et al. (2016) have recently redefined C. (Oecacta) with a synapomorphy and relegated a number of species previously placed in this subgenus or considered as miscellaneous to C. (Sensiculicoides). Nandi and Mazumdar (2014) suggested that the clavipalpis species group be placed within C. (Diphaomyia) but the

3 reasons for doing so are based on phenetic similarities that continue to plague a logical, cladistic basis for the classification of this genus. Some workers have placed single species as the sole representative of a species group (e.g. Khamala and Kettle, 1971) and I have simply included these in the list of miscellaneous species. As it stands now, species of Culicoides are placed in 32 subgenera and 38 species groups (unplaced to subgenus). Readers will see that there is a particularly long list of miscellaneous species, not placed in any group, representing 13% of the world fauna. A number of these, especially those in old publications, are likely nomina dubia. Unless types can be found, they need to be relegated to the dust bin by taxonomists - we already know that most of Kieffer's types are destroyed and most of his remaining names are therefore likely nomina dubia. The classification of African species is particularly messy and one need only compare the the following to see major discrepancies in the placement of many species: Khamala and Kettle (1971), Boorman and Dipeolu (1979), Itoua et al. (1987) and Glick (1990). At present there are no good keys to all the adults of species of any Region, although there are some local keys and/or guides that are very useful (these often only to females). A particularly innovative and excellent key for females of western Palaearctic species by Bruno Mathieu et al. is available online There are numerous further unnamed species from nearly every area on the planet. Large swaths, especially in tropical areas, have never been collected in any organized manner (e.g. most of the Andes and upper Amazon basin, Borneo). As we continue to explore the Earth's fauna, please, please, check types of previously described species from your own area as well as that of surrounding regions - we don't need more confusion in this arena either. There is a further important gap in our knowledge and application of some names of species of Culicoides because types of numbers of species need to be reexamined and studied carefully. A case in point is the current acceptance of Culicoides lupicaris as a synonym of C. delta in the catalog (which should only reflect the the present literature). Although evidence indicates that the two are actually valid and distinct species (Lassen et al., 2012;; Meiswinkel et al., 2004;; Ramilo et al., 2012, 2013), no one has formally made the change in a publication. The types should be reexamined to confirm the differences and ensure the names are properly applied. As such, there is a tremendous amount of systematic work that needs to be done. As the saying goes, "Use the following classification at your own risk!" If you have suggestions for improvements to the catalog below, please write to me at artborkent@telus.net

4 Full references for the taxonomic citations below are listed with the complete world catalog and are not repeated here. Some references above are not in the world catalog (no new names) and are given as follows: Ander, M., K. Troell, and J. Chirico Barcoding of biting midges in the genus Culicoides: a tool for species determination. Medical and Veterinary Entomology 27: Augot, D., F.J. Randrianambinintsoa, A. Gasser and J. Depaquit Record of two species of Culicoides (Diptera, Ceratopogonidae) new for Madagascar and molecular study showing the paraphylies of the subgenus Oecacta and the Schultzei group. Bulletin de la Societe de Pathologie Exotique 106: Bakhoum, M.T., M. Fall, A.G. Fall, G.A. Bellis, Y. Gottlieb, K. Labuschagne, G.J. Venter, M. Diop, I. Mall, M.T. Seck, X. Allene, M. Diarra, L. Gardes, J. Bouyer, J.-C. Delécolle, T. Balenghien, and C. Garros First record of Culicoides oxystoma Kieffer and diversity of species within the Schultzei Group of Culicoides Latreille (Diptera: Ceratopogonidae) Biting Midges in Senegal. PLOS ONE, 8 (12): /journal.pone Ekrem T, Willassen E, Stur E (2007) A comprehensive DNA sequence library is essential for identification with DNA barcodes. Molecular Phylogenetics and Evolution 43: doi: /j.ympev Ekrem T, Willassen E, Stur E (2010) Phylogenetic utility of five genes for dipteran phylogeny: A test case in the Chironomidae leads to generic synonymies. Molecular Phylogenetics and Evolution 57: doi: /j.ympev Itoua, A., M. Cornet, G. Vattier-Bernard, and J. Trouillet Les Culicoides (Diptera, Ceratopogonidae) de Afrique Centrale. Cahiers ORSTOM, Série Entomologie Médicale et Parasitologie, numéro spécial 1987: Lassen, S.B., S.A. Nielsen, H. Skovgard, and M. Kristensen Molecular differentiation of Culicoides biting midges (Diptera: Ceratopogonidae) from the subgenus Culicoides Latreille in Denmark. Parasitology Research 110: Linton, Y., A. Mordue-Luntz, R. Cruickshank, R. Meiswinkel, P. Mellor, and J. Dallas Phylogenetic analysis of the mitochondrial cytochromeoxidase subunit I gene of five species of the Culicoides imicola species complex. Medical and Veterinary Entomology. 16: Meiswinkel, R. and A.L. Dyce Afrotropical Culicoides: Synhelea Kieffer, 1925, resurrected as subgenus to embrace 10 species (Diptera: Ceratopogonidae). Onderstepoort Journal of Veterinary Research 56: Meiswinkel, R., L.M. Gomulski, J.-C. Delécolle, M. Goffredo, and G. Gasperi. 2004a. The taxonomy of Culicoides vector complexes - unfinished business. Vet. Ital. 40: Morag, N., Y. Saroya, Y. Braverman, E. Klement, and Y. Gottlieb, Y Molecular Identification, phylogenetic status, and geographic distribution of Culicoides

5 oxystoma (Diptera: Ceratopogonidae) in Israel. PLOS ONE, 7 (3): /journal.pone Ramilo, D.W., S. Diaz, I. Pereira da Fonseca, J.-C. Delécolle, A. Wilson, J. Meireles, J. Lucientes, R. Ribeiro, and F. Boinas First report of 13 species of Culicoides (Diptera: Ceratopogonidae) in mainland Portugal and Azores by morphological and molecular characterization. PLoS One 7(4):e Ramilo, D., C. Garros, B. Mathieu, C. Benedet, X. Allène, E. Silva, G. Alexandre-Pires, I.P. Da Fonseca, S. Carpenter, J. Rádrová and J.-C. Delécolle Description of Culicoides paradoxalis sp. nov. from France and Portugal (Diptera: Ceratopogonidae). Zootaxa 3745: Rivera J, Currie D (2009) Identification of Nearctic black flies using DNA barcodes (Diptera: Simuliidae). Molecular Ecology Resources 9: doi: /j x Schwenkenbecher, J.M., A.J. Mordue and S.B. Piertney Phylogenetic analysis indicates that Culicoides dewulfi should not be considered part of the Culicoides obsoletus complex. Bulletin of Entomological Research 99: Shults, P., A. Borkent and R. Gold The pupa of Culicoides sonorensis Wirth and Jones (Diptera: Ceratopogonidae) - first detailed description of this stage of the Bluetongue virus vector. Annals of the Entomological Society of America 109: ;; doi: /aesa/sav119 Szadziewski, R., S. Filatov and P. Dominiak A redescription of Culicoides griseidorsum Kieffer, 1918, with comments on subgeneric position of some European taxa (Diptera: Ceratopogonidae). Zootaxa 4107: Tay, W.T,, P.J. Kerr and L.S. Jermiin Population genetic structure and potential incursion pathways of the Bluetongue Virus vector Culicoides brevitarsis (Diptera: Ceratopogonidae) in Australia. PLOS ONE, 11 (1): /journal.pone

6 Genus CULICOIDES Latreille CULICOIDES Latreille, 1809: 251. Type species: Culicoides punctatus Latreille (= Ceratopogon punctatus Meigen), by monotypy. PADROSIA Rafinesque, 1815: 130 (unnecessary new name for Culicoides Latreille). Type species: Culicoides punctatus Latreille (= Ceratopogon punctatus Meigen), automatic. OXYHELEA Kieffer, 1921b: 14 (as subgenus of Culicoides). Type species: Culicoides dentatus Kieffer, by monotypy. PROSAPELMA Kieffer, 1925a: 417. Type species: Prosapelma cinerea Kieffer, by original designation. NEOCULICOIDES Boorman and Lane, 1979: 327 (preoccupied by Neoculicoides Pierce, 1966). Type species: Neoculicoides taylori Boorman and Lane, by original designation. DEVALQUIA Choufani and Nel, 2013: 75. Type species: Devalquia brisaci Choufani & Nel, 2013, by original designation. Subgenus AMOSSOVIA Glukhova AMOSSOVIA Glukhova, 1989: 226 (as subgenus of Culicoides). Type species: Culicoides dendrophilus Amosova, by original designation. arboricola Root and Hoffman, 1937: 166. USA (Maryland). beckae Wirth and Blanton, 1967: 213. USA (Florida). californiensis Wirth and Blanton, 1967: 215. USA (California). cochisensis Wirth and Blanton, 1967: 216. USA (Arizona). dendrophilus Amosova, 1957: 240. Russia. reesi Bullock and Akiyama, 1959: 24. Japan. elemae Pappas and Pappas, 1989: 228. USA (Nebraska). flukei Jones, 1956: 30. USA (Wisconsin). guttipennis (Coquillett), 1901: 603 (Ceratopogon). USA (Ohio). oklahomensis Khalaf, 1952b: 355 (as villosipennis spp.). USA (Oklahoma). ousairani Khalaf, 1952b: 354. USA (Oklahoma). pecosensis Wirth, 1955: 358. USA (Texas). villosipennis Root and Hoffman, 1937: 165. USA (Maryland). Subgenus ANILOMYIA Vargas ANILOMYIA Vargas, 1960: 37 (as subgenus of Culicoides). Type species: Culicoides covagarciai Ortiz, by original designation. ameliae Browne, 1980: 543. Colombia. chaverrii Spinelli and Borkent, 2004a: 373. Costa Rica. chrysonotus Wirth and Blanton, 1956b: 226. Panama. covagarciai Ortiz, 1950b: 457. Venezuela. beebei Fox, 1952: 366. Venezuela. decor (Williston), 1896: 281 (Ceratopogon). St. Vincent. dominicanus Wirth and Blanton, 1970d: 146. Dominica. efferus Fox, 1952: 365. Peru. farri Wirth and Blanton, 1970d: 148. Jamaica. hayesi Matta, 1967: 75. Honduras. inermis Spinelli, Santamaría, Cabrera, Ronderos and Suárez, 2009: 84. Colombia. lutealaris Wirth and Blanton, 1956b: 225. Panama. marshi Wirth and Blanton, 1956b: 220. Panama. metagonatus Wirth and Blanton, 1956b: 221. Panama.

7 monicae Spinelli and Borkent, 2004a: 372. Costa Rica. nigrifemur Spinelli, Santamaría, Cabrera, Ronderos and Suárez, 2009: 85. Colombia. nigrigenus Wirth and Blanton, 1956b: 222. Panama. popayanensis Wirth and Lee, 1967: 10. Colombia. pseudodecor Spinelli and Huerta, 2015: 818. Mexico. rostratus Wirth and Blanton, 1956b: 218. Panama. trapidoi Wirth and Barreto, 1978: 554. Colombia. Subgenus AVARITIA Fox AVARITIA Fox, 1955: 218 (as subgenus of Culicoides). Type species: Ceratopogon obsoletus Meigen, by original designation. abchazicus Dzhafarov, 1964: 263. Georgia. actoni Smith, 1929: 255 (1932: 180). India. okumensis Arnaud, 1956a: 119. Japan. imperceptus Das Gupta, 1962a: 538. India. alachua Jamnback and Wirth, 1963: 187. USA (Florida). albifascia Tokunaga, 1937a: 319. Taiwan. alticola Kieffer, 1913e: 11. Tanzania. andicola Wirth and Lee, 1967: 5. Colombia. annandalei Majumdar and Das Gupta, in Gangopadhyay and Das Gupta, 2000: 125. India. asiana Bellis, in Bellis et al. 2015: 29. New name for asiatica. asiatica Bellis, in Bellis et al. 2013: 407 (preoccupied by Culicoides asiaticus Gutsevich and Smatov, 1966). Japan. autumnalis Sen and Das Gupta, 1959a: 628. India. bolitinos Meiswinkel, 1989: 30. South Africa. boophagus Macfie, 1937c: 116. Malaysia. boydi Wirth and Mullens, 1992: USA (California). brevipalpis Delfinado, 1961: 654. Philippines. brevitarsis Kieffer, 1917a: 187. Australia. robertsi Lee and Reye, 1953: 386. Australia (Queensland). radicitus Delfinado, 1961: 657. Philippines. superfulvus Das Gupta, 1962b: 253. India. brosseti Vattier and Adam, 1966a: 297. Gabon. bubalus Delfinado, 1961: 658. Philippines. candolfii Delécolle, Paupy, Rahola, and Mathieu, 2013: 514. Gabon. certus Das Gupta, 1962a: 537. India. chiopterus (Meigen), 1830: 263 (Ceratopogon). Europe. amoenus (Winnertz), 1852: 35 (Ceratopogon). Germany. similis (Goetghebuer), 1927a: 203 (Monohelea, preoccupied by Culicoides similis Carter, Ingram and Macfie, 1920). Belgium. dobyi Callot and Kremer, 1969a: 610. France. comparis Liu and Yu, in Yu et al., 2006: 917. China. conaensis Liu and Yu, 1990: 19. China. dasyops Clastrier, 1958b: 217. Senegal. definitus Sen and Das Gupta, 1959a: 622. India. dentiformis McDonald and Lu, 1972: 403. Taiwan. dewulfi Goetghebuer, 1936: 320. Belgium. pseudochiopterus Downes and Kettle, 1952: 67. Great Britain. dikhros Tokunaga, 1962b: 507. Papua New Guinea. dubitatus Kremer, Rebholtz-Hirtzel and Delécolle, 1976: 233. Angola. dumdumi Sen and Das Gupta, 1959a: 628. India. elongatus Chu and Liu, 1978: 83. China.

8 filicinus Gornostaeva and Gachegova, 1972: 522. Russia. flavipunctatus Kitaoka, 1975: 199. Japan. fragmentum Tokunaga, 1962b: 507. Papua New Guinea. fulvus Sen and Das Gupta, 1959a: 628. India. gaponus Yu, 1982: 202. China. glabripennis Goetghebuer, 1935d: 171 (1948b: 12). New name for nudipennis. nudipennis Goetghebuer, 1933e: 146 (preoccupied by Culicoides nudipennis Kieffer, 1922g). Democratic Republic of Congo. septemmaculatus Goetghebuer, 1935d: 175. Democratic Republic of the Congo. spinifer Khamala and Kettle, 1971: 51. Kenya. glushchenkoae Glukhova, 1989: 200. Russia gornostaevae Mirzaeva, 1984a: 371. Russia. grahamii Austen, 1909: 280. Ghana. habereri Becker, 1909: 289. Cameroon. hostilissimus (Pittaluga), 1911a: 29 (1911b: 69;; 1912: 597) (Oecacta). Guinea. kivuensis Goetghebuer, 1935d: 173. Democratic Republic of the Congo. trichopis de Meillon, 1937b: 328. Democratic Republic of the Congo. gulbenkiani Caeiro, 1959: 155. South Africa. hasegawai Kanasugi and Kitaoka, 2001:228. Japan. hayakawai Kitaoka, 1984: 301. Japan. hermani Spinelli and Borkent, 2004a: 363. Panama. himalayae Kieffer, 1911b: 326. India. hirtulus (Coquillett), 1900: 396 (Ceratopogon). USA (Alaska). holcus Lee, 1980: 85. China. huambensis Caeiro, 1961: 179 (also as huambenis). Angola. hui Wirth and Hubert, 1961: 16. Taiwan. imicola Kieffer, 1913e: 11. Kenya. pallidipennis Carter, Ingram and Macfie, 1920: 265. Ghana. iraqensis Khalaf, 1957b: 343. Iraq. minutus Sen and Das Gupta, 1959a: 622. India. pseudoturgidus Das Gupta, 1962a: 537. India. impusilloides Spinelli and Wirth, 1984a: 178. Brazil. incertus Yu and Zhang, in Yu, 1988: 136. China. inexploratus Sen and Das Gupta, 1959a: 628. India. insignipennis Macfie, 1937d: 469. Malaysia. iphthimus Zhou and Lee, 1984b: 295. China. jacobsoni Macfie, 1934d: 215. Indonesia. buckleyi Macfie, 1937c: 117. Malaysia. kitaokai Tokunaga, 1955: 6. Japan. unisetiferus Tokunaga, 1959: 236. Papua New Guinea. juddi Cochrane, 1973: 316. USA (New York). kanagai Khamala and Kettle, 1971: 49. Kenya. kibatiensis Goetghebuer, 1935d: 172. Democratic Republic of the Congo. volatilis Goetghebuer, 1935d: 176. Democratic Republic of the Congo. lengi Yu and Liu, 1990: 10. China. liweiae Liang, Fu and Liu, 2012: 234. China. longirostris Qu and Wang, 1994: 486. China. loxodontis Meiswinkel, 1992: 147. South Africa. mamaensis Lee, 1979a: 101. China. minimus Wirth and Hubert, 1989: 251. Malaysia. miombo Meiswinkel, 1991: 161. Malawi. molestior Kieffer, 1911e: 514. New name for molestus. molestus Kieffer, 1910: 192 (preoccupied by Culicoides molestus (Skuse), 1889). India. iniquus Sen and Das Gupta, 1958b: 163 (1959a: 620;; 1959b: 65). New name for molestus. montanus Shakirzjanova, 1962: 258. Kazakhstan.

9 motoensis Lee, 1978: 75 (1979a: 102). China. nielamensis Liu and Deng, 2000: 246. China. nigritus Fei and Lee, 1984b: 345. China. nudipalpis Delfinado, 1961: 655. Philippines. nujiangensis Liu, 1990: 59. China. nupurius Kanasugi and Kitaoka, 2001:227. Japan. obscurus Tokunaga and Murachi, 1959: 347. Belau (USA). pungens de Meijere, 1909: 200 (preoccupied by Culicoides pungens (Kieffer), 1901a). Indonesia. obsoletus (Meigen), 1818: 76 (Ceratopogon). Europe. varius (Winnertz), 1852: 35 (Ceratopogon). Germany. yezoensis (Matsumura), 1911: 60 (Ceratopogon). Russia. obscuripes Santos Abreu, 1918: 297 (as variety of varius). Canary Islands (Spain). lacteinervis Kieffer, 1919a: 47. Slovak Republic, Ukraine. rivicola Kieffer, 1921a: 56. Germany. clavatus Kieffer, 1921a: 56. Germany. heterocerus Kieffer, 1921a: 57. Germany. pegobius Kieffer, 1922c: 235. Germany. kabyliensis Kieffer, 1922g: 505. Algeria. concitus Kieffer, 1922e: 71. Germany. intermedius Okada, 1941: 22 (as variety of obsoletus, preoccupied by Dasyhelea intermedia (Santos Abreu), 1918). Japan. sintrensis Cambournac, 1956: 591. Portugal. seimi Shevchenko, 1967: 173. Ukraine. odiosus Kieffer, 1910: 192. India. orientalis Macfie, 1932b: 490. Malaysia. nayabazari Das Gupta, 1963: 35. India. orjuelai Wirth and Lee, 1967: 6. Colombia. palauensis Tokunaga, in Tokunaga and Murachi, 1959: 348. Belau (USA). pastus Kitaoka, 1980: 11. Japan. pechumani Cochrane, 1974: 133. USA (New York). pelius Liu and Yu, 1990: 23. China. pseudopallidipennis Clastrier, 1958b: 197. Senegal. puracensis Wirth and Lee, 1967: 7. Colombia. pusilloides Wirth and Blanton, 1955a: 104. Panama. pusillus Lutz, 1913: 52. Brazil. qionghaiensis Yu and Liu, 1990: 4. China. ruiliensis Lee, 1980: 86. China. sanguisuga (Coquillett), 1901a: 604 (Ceratopogon). USA (Maryland). scoticus Downes and Kettle, 1952: 65. Great Britain. sikkimensis Das Gupta, 1963: 35. India. sinanoensis Tokunaga, 1937a: 331. Japan. obsoletiformis Amosova, 1957: 233. Russia. sousadiasi Caeiro, 1961: 309. Angola. suarezi Rodriguez and Wirth, 1986: 313. Colombia. suiyangensis Hou, Han, Lv and Jiang, China. suzukii Kitaoka, 1973: 212. Japan. tainanus Kieffer, 1916b: 114. Taiwan. maculatus (Shiraki), 1913: 294 (Ceratopogon, preoccupied by Atrichopogon maculatus (Lundström), 1910). Taiwan. kii Tokunaga, 1937a: 284. Japan. sigaensis Tokunaga, 1937a: 322. Japan. kyotoensis Tokunaga, 1937a: 329. Japan. suborientalis Tokunaga, 1951: 106. Indonesia. tibetensis Chu, 1977: 102. China. tororoensis Khamala and Kettle, 1971: 48. Uganda.

10 trifasciellus Goetghebuer, 1935d: 175. Democratic Republic of the Congo. trimaculatus McDonald and Lu, 1972: 415. Taiwan. tuttifrutti Meiswinkel, Cornet and Dyce, in Meiswinkel and Linton, 2003: 42. South Africa. wadai Kitaoka, 1980: 14. Japan. wandashanensis Wang and Liu, 1999: 328. China. yamii Lien, Lin and Weng, 1998b:57. Taiwan. yuchihensis Lien, Lin and Weng, 1998b:58. Taiwan. Subgenus BELTRANMYIA Vargas BELTRANMYIA Vargas, 1953: 34 (as subgenus of Culicoides). Type species: Culicoides crepuscularis Malloch, by original designation. alaskensis Wirth, 1951a: 84. USA (Alaska). bermudensis Williams, 1956: 298. Bermuda (Great Britain). chagyabensis Lee, 1982: 169. China. charadraeus Arnaud, 1956a: 97. Japan. chengduensis Zhou and Lee, 1984a: 222. China. circumscriptus Kieffer, 1918a: 49. Tunisia. nadayanus Kieffer, 1918a: 95. Turkey. edwardsi Goetghebuer, 1921: 177. Belgium. algarum Kieffer, 1924a: 18. Germany. salicola Kieffer, 1924b: 405. Norway. pictidorsum Kieffer, 1924b: 406 (as variety of salicola). Norway. albonotatus Vimmer, 1932: 133 (preoccupied by Culicoides albonotatus Kieffer, 1918a). Israel. albosignatus Vimmer, 1932: 135. Israel. polymaculatus Vimmer, 1932: 135. Israel. pulcher Zilahi-Sebess, 1934: 155 (misspelled as pulscher). Bulgaria. kirovabadicus Dzhafarov 1964: 228. Azerbaijan. matsuensis Lien, Weng and Lin, 1996: 20 (valid subspecies of circumscriptus). Taiwan. meridionalis Xue, Liu and Yu, 2003: 112 (valid subspecies of circumscriptus). China. corniculus Liu and Qu, 1981: 419. China. apiculatus Yu and Zhang, in Yu, 1988: 134. China. crepuscularis Malloch, 1915a: 303. USA (Illinois). desertorum Gutsevich, 1959: 675. Turkmenistan and Uzbekistan. pictus Khalaf, 1961: 455. Iraq. brevifrontis Smatov and Isimbekov, 1971: 62. Kazakhstan. oxianus Smatov, in Smatov and Kravets, 1976: 282. Kazakhstan. duodenarius Kieffer, 1922b: 157. Taiwan. gluchovae Mirzaeva, 1974: 27. Russia. wushenensis Lee (as Li), 1974: 353. China. halonostictus Wirth and Hubert, 1989: 431. Thailand. hollensis (Melander and Brues), 1903: 13 (Ceratopogon). USA (Massachusetts). canithorax Hoffman, 1925: 284. USA (Georgia). homochrous Remm, in Remm and Zhogolev, 1968: 835. Ukraine. huayingensis Zhou and Lee, 1984b: 293. China. japonicus Arnaud, 1956a: 106. Japan. litoreus Amosova, 1957: 241. Russia. knowltoni Beck, 1956: 136. USA (Florida). koreensis Arnaud, 1956a: 109. South Korea. palustris Amosova, 1957: 243. Russia. liubaensis Liu and Wu, 2005:448. China. manchuriensis Tokunaga, 1941b: 98. China. setiger Goetghebuer, 1938: 379 (preoccupied by Dasyhelea setigera (Kieffer), 1910). Belgium. goetghebueri Arnaud, 1956b: 94. Belgium. New name for setiger. machardyi Campbell and Pelham-Clinton, 1960: 235. Great Britain.

11 ochraceimaculatus Shevchenko, 1970b: 8. Ukraine. ochraceipennis Shevchenko, 1970b: 9. Ukraine. mesostigma Remm, 1971: 201. Russia. vistulensis Skierska, 1973: 289. Poland. mississippiensis Hoffman, 1926a: 158. USA (Mississippi). multifidous Liu and Yu, in Yu et al., 2006: China. musajevi Dzhafarov, 1961c: 75. Azerbaijan. navaiae Lane, 1983: 534. Saudi Arabia. nivosus de Meillon, 1937b: 341. South Africa. pallidulus Yu, 1991: 52. New name for obscuratus. obscuratus Ding and Yu, 1990: 54 (preoccupied by Culicoides obscuratus Statz, 1944). China. pseudosalinarius Chu, 1981: 309. China. pulchellus Liu and Zhao, 1998: 12. China. qabdoensis Lee, 1979a: 103. China. salinarius Kieffer, 1914a: 236. Germany. halobius Kieffer, 1914a: 237. Germany. meinerti Kieffer, 1915c: 284. Denmark. punctatidorsum Kieffer, 1924a: 17. Germany. shanghaiensis Qu, Cao and Liu, 2010: 322. China. sibiricus Mirzaeva, 1964: 218. Russia. sphagnumensis Williams, 1955: 269. USA (Michigan). lacicola Arnaud, 1956a: 110 (as laciocola). Japan. carjalaensis Glukhova, 1957: 249. Russia. stellaris Yu and Liu, 1990: 6. China. subarakawae Yu and Zou, in Hao et al., 1990: 41. China. subcircumscriptus Yu, 1982: 202. China. superflutheca Yu and Li, in Yu et al., 1986: 210. China. suspectus Zhou and Lee, 1984b: 294. China. tessellatus Yu, 1982: 202. China. toyamaruae Arnaud, 1956a: 133. Japan. wisconsinensis Jones, 1956: 32. USA (Wisconsin). xenkeensis Yang, Li and Liu, 2011: 154. China. yigongensis Liu and Deng, 2010: 578. China. Subgenus COTOCRIPUS Brèthes COTOCRIPUS Brèthes, 1912: 451. Type species: Cotocripus caridei Brèthes, by monotypy. bambusicola Lutz, 1913: 62. Brazil. bahiensis Barbosa, 1947: 11. Brazil. caridei (Brèthes), 1912: 451 (Cotocripus). Argentina. setifer (Lutz), 1913: 64 (Centrorhynchus). Brazil. gabrieli Spinelli, Santamaría, Cabrera, Ronderos and Suárez, 2009: 82. Colombia. irwini Spinelli and Wirth, 1984a: 180. Chile. patagoniensis Ronderos and Spinelli, 1997: 34. Argentina. raposoensis Wirth and Barreto, 1978: 561. Colombia. Subgenus CULICOIDES Latreille SILVICOLA Mirzaeva and Isaev, 1990: 98 (as subgenus of Culicoides). Type species: Culicoides grisescens Edwards, by original designation. almeidae Cambournac, 1970b: 251. Portugal. angkaensis Kitaoka, Takaoka and Choochote, 2005: 285. Thailand. aomoriensis Kitaoka, 1991: 289. Japan.

12 aterinervis Tokunaga, 1937a: 312. Japan. boyi Nielsen, Kristensen and Pape, 2015: 3. Denmark. brucei Austen, 1909: 282. Uganda. pseudopulicaris Goetghebuer, 1935d: 173. Democratic Republic of the Congo. hirtius de Meillon and Lavoipierre, 1944: 58. South Africa. choochotei Kitaoka and Takaoka, in Kitaoka et al, 2005: 287. Thailand. cockerellii (Coquillett), 1901: 603 (Ceratopogon). USA (Colorado). delta Edwards, in Edwards et al., 1939: 48. Great Britain. lupicaris Downes and Kettle, 1952: 76. Great Britain. dubius Arnaud, 1956a: 100. Japan. elutus Macfie, 1948: 75. Mexico. fagineus Edwards, in Edwards et al., 1939: 147. Great Britain. flavipulicaris Dzhafarov, 1964: 238. Azerbaijan. flavus Gornostaeva, 1980: 81 (as subspecies of grisescens). Russia. flavisomum Mirzaeva, 1984b: 67. Russia. anadyriensis Mirzaeva, 1984b: 70. Russia. fortinensis Spinelli and Huerta, 2015: 812. Mexico. freeborni Wirth and Blanton, 1969b: 217. USA (California). frohnei Wirth and Blanton, 1969b: 219. USA (Alaska). gregsoni Wirth and Blanton, 1969b: 222. Canada (British Columbia). grisescens Edwards, in Edwards et al., 1939: 146. Great Britain. remmi Damian-Georgescu, 1972: 16. Romania. arschanicus Mirzaeva, 1984b: 69. Russia. hondurensis Spinelli and Borkent, 2004a: 369. Honduras. hulinensis Liu and Yu, 1996: 135 (Liu et al., 1999: 341). China. impunctatus Goetghebuer, 1920: 55. Belgium. minor Tokunaga, 1941b: 97 (as variety of impunctatus). China. kalix Nielsen, Kristensen and Pape, 2015: 9. Sweden. kelinensis Lee, 1978: 58 (1979a: 99). China. kirinensis Lee, 1976: 47. China. lahontan Wirth and Blanton, 1969b: 223. USA (California). lanyuensis Kitaoka and Tanaka, 1985: 39. Taiwan. lungchiensis Chen and Tsai, 1962: 397 (as subspecies of peregrinus). China. megaforticeps Kitaoka, 1973: 212. Japan. luteovenus Root and Hoffman, 1937: 156. Mexico. magnus Colaco, 1946: 219, 237 (as variety of hirtius). South Africa. mathisi Giles and Wirth, 1983: 36. Sri Lanka. mcdonaldi Wirth and Hubert, 1989: 224. New name for monticola. monticola McDonald and Lu, 1972: 413 (as subspecies of pulicaris). Taiwan. neofagineus Wirth and Blanton, 1969b: 227. USA (Arizona). neomontanus Wirth, 1976b: 15 (new name for montanus Wirth and Blanton). USA (Utah). montanus Wirth and Blanton, 1969b: 225 (preoccupied by Shakirzjanova, 1962). USA (Utah). neopulicaris Wirth, 1955: 355. USA (Texas). newsteadi Austen, 1921: 113. Israel. biclavatus Kieffer, 1924a: 14. Germany. halophilus Kieffer, 1924b: 404. Norway. edwardsi Goetghebuer, 1933f: 46 (as variety of pulicaris, preoccupied by Culicoides edwardsi Goetghebuer, 1921). Europe. edwardsianus Goetghebuer, 1933d: 367 (as variety of pulicaris). Great Britain. nipponensis Tokunaga, 1955: 4. Japan. padusae Mirzaeva, 1989: 87 (1990: 90). Russia. papilliger Borkent, in Borkent and Wirth, 1997:78. New name for papillatus. papillatus Kitaoka and Shinonaga, 1989: 26 (preoccupied by Culicoides papillatus Khamala and Kettle, 1971). Pakistan.

13 paradoxalis Ramilo and Delécolle, in Ramilo et al., 2013: 244. France. paraimpunctatus Borkent, 1995: 60. USA (Minnesota). canadensis Wirth and Blanton, 1969b: 211 (preoccupied by Boesel, 1937). USA (Minnesota). pulicaris (Linnaeus), 1758: 603 (Culex). Europe. setosinervis Kieffer, 1913a: 8. Germany. pullatus Kieffer, 1915a: 474. Germany. stephensi Carter, 1916: 135. Egypt. cinerellus Kieffer, 1919a: 40. Ukraine. quinquepunctatus Goetghebuer, 1921: 177. Belgium. flaviplumus Kieffer, 1924a: 19. Germany. sawamotoi Kono and Takahasi, 1940: 75. Russia. punctatus (Meigen), 1804: 29 (Ceratopogon). Europe. punctatus Latreille, 1809: 252 (preoccupied by Culicoides punctatus (Meigen), 1804). France. ocellaris Kieffer, 1921d: 276 (as variety of pulicaris). Latvia. kasachstanicus Shakirzjanova, 1963: 63 (as subspecies of pulicaris). Kazakhstan. putianensis Lee, 1988: 96. China. qufuensis Xue and Yu, in Xue et al., 1992: 52. China. quqiaoensis Chen, in Lee, 1988: 99. China. rulfoi Spinelli and Huerta, 2015: 816. Mexico. saltonensis Wirth, 1952a: 173 (as cockerellii ssp.). USA (California). selandicus Nielsen, Kristensen and Pape, 2015: 6. Denmark. sellersi Boorman and Dipeolu, 1979: 55. Nigeria. sierrensis Wirth and Blanton, 1969b: 232. USA (California). sommermanae Wirth and Blanton, 1969b: 234. USA (Alaska). sordidellus (Zetterstedt), 1838: 820 (Ceratopogon). Greenland (Denmark). subfagineus Delécolle and Ortega, 1998: 285. Spain. subpunctatus Liu and Yu, 1996: 137 (Liu et al., 1999: 341). China. tahemanensis Liu and Ma, 2001: 199. China. tayulingensis Chen, 1983: 77 (1988: 153). China. tienhsiangensis Chen, 1983: 78 (1988: 153). China. tristriatulus Hoffman, 1925: 294 (as cockerelli var.). USA (California). yukonensis Hoffman, 1925: 291. Canada (Yukon Territory). yunanensis Chu and Liu, 1978: 81. China. Subgenus DIPHAOMYIA Vargas DIPHAOMYIA Vargas, 1960: 40 (as subgenus of Culicoides). Type species: Culicoides baueri Hoffman, by original designation. baueri Hoffman, 1925: 297. USA (Maryland). bergi Cochrane, 1973: 311. USA (New York). blantoni Vargas and Wirth, 1955: 33. Mexico. defoliarti Atchley and Wirth, 1979: 527. USA (Arizona). edeni Wirth and Blanton, 1974b: 23. USA (Florida). erikae Atchley and Wirth, 1979: 532. USA (New Mexico). evansi Wirth and Blanton, 1959: 342. Panama. footei Wirth and Jones, 1956: 162. USA (Virginia). freitasi Wirth and Blanton, 1973: 434. Brazil (Para). haematopotus Malloch, 1915a: 302. USA (Illinois). inyoensis Wirth and Blanton, 1969a: 565. USA (California). iriartei Fox, 1952: 368. Venezuela. vargasi Wirth and Blanton, 1953a: 74. Panama. jurbergi Felippe-Bauer, in Felippe-Bauer, et al., 2005: 51. Peru. lisicarruni Moncada, Carrasquilla and Spinelli, 2010: 978. Colombia. marinkellei Wirth and Lee, 1967: 13. Colombia. mckeeveri Brickle and Hagan, 1999: 39. Belize.

14 minasensis Felippe-Bauer, 1987: 147. Brazil. mirsae Ortiz, 1953a: 801. Venezuela. mukerjii Majumdar and Das Gupta, in Gangopadhyay and Das Gupta, 2000: 125. India. peculiaris Majumdar and Das Gupta, in Gangopadhyay and Das Gupta, 2000: 127. India. ronderosae Spinelli and Borkent, 2004a: 377. Costa Rica. soleamaculatus Nandi and Mazumdar, 2014: 468. India. tarapaca Spinelli and Wirth, 1984a: 184. Chile. Subgenus DRYMODESMYIA Vargas DRYMODESMYIA Vargas, 1960: 40 (as subgenus of Culicoides). Type species: Culicoides copiosus Root and Hoffman, by original designation. arizonensis Wirth and Hubert, 1960: 655. USA (Arizona). bakeri Vargas, 1954: 27. Mexico. borinqueni Fox and Hoffman, 1944: 110. Puerto Rico (USA). bredini Wirth and Blanton, 1970b: 41. Dominica. butleri Wirth and Hubert, 1960: 650. USA (Arizona). byersi Atchley, 1967: 983. USA (New Mexico). cacticola Wirth and Hubert, 1960: 653. USA (California). chacoensis Spinelli and Wirth, 1984a: 174. Argentina. copiosus Root and Hoffman, 1937: 171. Mexico. haitiensis Delécolle, Raccurt and Rebholtz, 1986: 108. Haiti. hinmani Khalaf, 1952b: 353. USA (Oklahoma). hitchcocki Spinelli and Wirth, 1984a: 176. Peru. insolatus Wirth and Hubert, 1960: 654. Mexico. jamaicensis Edwards, 1922: 165 (as loughnani var.). Jamaica. jonesi Wirth and Hubert, 1960: 650. USA (Texas). loughnani Edwards, 1922: 165. Jamaica. panamensis Barbosa, 1947: 22. Panama. alambiculorum Macfie, 1948: 81. Mexico. pilosus Wirth and Blanton, 1959: 332. Panama. poikilonotus Macfie, 1948: 82. Mexico. cacozelus Macfie, 1948: 85. Mexico. hertigi Wirth and Blanton, 1953b: 229. Panama. ryckmani Wirth and Hubert, 1960: 656. USA (California). saltaensis Spinelli and Wirth, 1984a: 183. Argentina. sitiens Wirth and Hubert, 1960: 652. USA (California). torridus Wirth and Hubert, 1960: 654. Mexico uruguayensis Ronderos, 1990a: 117. Uruguay. wirthomyia Vargas, 1953b: 227. Mexico. Subgenus FASTUS Liu FASTUS Liu, in Yu et al., 2006: 1196 (as subgenus of Culicoides). Type species: Culicoides alpigenus Yu and Liu, by original designation. alpigenus Yu and Liu, in Yu et al., 2006: China. changbaiensis Qu and Ye, 1995:95. China. dispersus Gutsevich and Smatov, 1966: 76. Kazakhstan. erairai Kono and Takahasi, 1940: 70. Japan. haerbalingensis Liu and Wang, in Liu et al., 2010: 63. China. huochengensis Ma and Yu, in Ma et al., 1990: 47. China. kangdingensis Yu and Liu, in Yu et al., 2006: China. pingtanensis Yu, 1982: 202. China.

15 subchangbaiensis Liu and Ren, in Liu et al., 2010: 64. China. taipingshanensis Liu and Guo, 2009:50. China. zyzzamaculatus Yu, 1982: 202. China. Subgenus GLAPHIROMYIA Vargas GLAPHIROMYIA Vargas, 1960: 41 (as subgenus of Culicoides). Type species: Culicoides scopus Root and Hoffman, by original designation. dampfi Root and Hoffman, 1937: 169. Mexico. parascopus Wirth and Blanton, 1978: 238. Mexico. scopus Root and Hoffman, 1937: 170. Mexico. Subgenus HAEMOPHORUCTUS Macfie HAEMOPHORUCTUS Macfie, 1925: 349. Type species: Haemophoructus maculipennis Macfie, by monotypy. boormani Giles and Wirth, 1985: 365. Malaysia. calcaratus Wirth and Hubert, 1989: 157. Malaysia. gemellus Macfie, 1934c: 192. Malaysia. gentilis Macfie, 1934c: 191. Malaysia. gentiloides Kitaoka and Tanaka, 1985: 41. Taiwan. gymnopterus Edwards, 1926b: 247. Malaysia. hoffmanioides Wirth and Hubert, 1989: 173. Malaysia. kinari Howarth, 1985: 44. Laos. kisangkini Howarth, 1985: 47. Laos. maculipennis (Macfie), 1925: 349 (Haemophoructus). Singapore. mellipes Wirth and Hubert, 1989: 179. Malaysia. nitens Edwards, 1933c: 252. Malaysia. nyakini Howarth, 1985: 49. Laos. rariradialis Das Gupta, 1963: 34. India. tawauensis Wirth and Hubert, 1989: 161. Malaysia. unicus Delfinado, 1961: 665. Philippines. Subgenus HAEMATOMYIDIUM Goeldi HAEMATOMYIDIUM Goeldi, 1905: 137. Type species: Haematomyidium paraensis Goeldi, by original designation. annuliductus Wirth, in Vitale et al., 1981: 150. Panama. aragaoi Tavares and Luna Dias, 1980: 393. Brazil. austroparaensis Spinelli, in Spinelli et al., 2005: 141. Argentina. bayano Wirth, in Vitale et al., 1981: 152. Panama. crucifer Clastrier, 1968: 85. French Guiana. darlingtonae Wirth and Blanton, 1971a: 39. Trinidad and Tobago. debilipalpis Lutz, 1913: 60. Brazil. khalafi Beck, 1957: 104. USA (Florida). ichesi Ronderos and Spinelli, 1995b: 77. Argentina. denisae Clastrier, 1971: 290. French Guiana. diversus Felippe-Bauer, in Felippe-Bauer, et al., 2003: Peru. dureti Ronderos and Spinelli, 1995a: 59. Paraguay. eadsi Wirth and Blanton, 1971a: 37. USA (Texas). eldridgei Wirth and Barreto, 1978: 561. Colombia. equatoriensis Barbosa, 1952: 13 (1953: 14) (as variety of debilipalpis). Ecuador. espinolai Felippe-Bauer and Lourenco-de-Oliveira, 1987: 149. Brazil.

16 filiductus Wirth, in Vitale et al., 1981: 155. Panama. flinti Wirth, 1982b: 251. Argentina. germanus Macfie, 1940d: 27. Guyana. ginesi Ortiz, 1951c: 586. Venezuela. glabrior Macfie, 1940d: 27 (as variety of debilipalpis). Guyana. grahambelli Forattini, 1956b: 35. Panama. hoffmani Fox, 1946a: 251. Trinidad and Tobago. horticola Lutz, 1913: 61. Brazil. bachmanni Spinelli, in Spinelli et al., 2005: 146. Argentina. imitator Ortiz, 1953b: 808. Venezuela. insinuatus Ortiz and León, 1955: 577. Ecuador. jurutiensis Trindade and Felippe-Bauer, 2011a:61. Brazil. kampa Felippe-Bauer, Veras, Castellon and Moreira, 2000: 35. Brazil. kettlei Breidenbaugh and Mullens, 1999: 150. USA (California). lahillei (Iches), 1906: 264 (Ceratopogon). Argentina. limonensis Ortiz and León, 1955: 576. Ecuador. martyrius Trindade and Felippe-Bauer, 2011a:62. Brazil. neoparaensis Tavares and Souza, 1978: 621. Brazil. pampaensis Spinelli and Wirth, 1984a: 182. Argentina. paraensis (Goeldi), 1905: 137 (Haematomyidium). Brazil. undecimpunctatus Kieffer, 1917b: 307. Argentina. peruvianus Felippe-Bauer, in Felippe-Bauer, et al., 2003: Peru. quasiparaensis Clastrier, 1971: 286. French Guiana. rachoui Tavares and Souza, 1978: 622. Brazil. spurius Wirth and Blanton, 1959: 433. Panama. todatangae Wirth and Blanton, 1973: 447. Brazil. torreyae Wirth and Blanton, 1971c: 73. USA (Florida). youngi Wirth and Barreto, 1978: 562. Colombia. Subgenus HOFFMANIA Fox HOFFMANIA Fox, 1948: 21 (as subgenus of Culicoides). Type species: Culicoides inamollae Fox and Hoffman (= Culicoides insignis Lutz), by original designation. aitkeni Wirth and Blanton, 1968a: 214. Trinidad and Tobago. andrewsi Causey, 1938: 410. Thailand. annettae Spinelli and Borkent, 2004a: 365. Costa Rica. antioquiensis Spinelli, Santamaría, Cabrera, Ronderos and Suárez, 2009: 82. Colombia. baniwa Felippe-Bauer, in Felippe-Bauer, et al., 2009: 852. Brazil. batesi Wirth and Blanton, 1973: 426. Brazil. sanmartini Wirth and Barreto, 1978: 553. Colombia. bawanglingensis Yu,Wang and Chen, in Wang, et al., 2012: 283. China. biestroi Spinelli and Ronderos, 1991: 86. Argentina. bimaculatus Floch and Abonnenc, 1942b: 3. French Guiana. brasilianum Forattini, 1956a: 81. Brazil. brinchangensis Wirth and Hubert, 1989: 190. Malaysia. brownei Spinelli, in Spinelli et al., 1993: 24. Colombia. calchaqui Spinelli and Veggiani Aybar, in Spinelli, et al., 2013: 586. Argentina. cameronensis Kitaoka, 1983: 94. Malaysia. carpophilus Wirth and Hubert, 1989: 196. Malaysia. charruus Spinelli and Martinez, 1992: 176. Uruguay. cheahi Kitaoka, 1983: 92. Malaysia. contubernalis Ortiz and León, 1955: 574 (as variety of rozeboomi). Ecuador. coutinhoi Barretto, 1944: 96. Brazil. davidi Spinelli, in Spinelli et al., 1993: 30. Colombia. diabolicus Hoffman, 1925: 294. Panama.

17 diffusus Spinelli, in Spinelli et al., 1993: 34. Brazil. divisus Wirth and Hubert, 1989: 198. Malaysia. effusus Delfinado, 1961: 658. Philippines. fernandoi Tavares and Sousa, 1979: 611. Brazil. ferreyrai Ronderos and Spinelli, 1995a: 61. Argentina. filariferus Hoffman, 1939: 172. Mexico. flavivenulus Costa Lima, 1937a: 418. Brazil. foxi Ortiz, 1950c: 461. Puerto Rico (USA). franklini Spinelli, in Spinelli et al., 1993: 45. Panama. fusipalpis Wirth and Blanton, 1973: 435. Brazil. guttatus (Coquillett), 1904c: 35 (Ceratopogon). Brazil. heliconiae Fox and Hoffman, 1944: 108. Venezuela. rozeboomi Barbosa, 1947: 26. Trinidad and Tobago. hirtipennis Delfinado, 1961: 662. Philippines. hylas Macfie, 1940d: 26. Guyana. ignacioi Forattini, 1957: 215. Brazil. saintjusti Tavares and Ruiz, 1980: 27. Brazil. indianus Macfie, 1932b: 488. India. cylindratus Kitaoka, 1980: 16. Japan. innoxius Sen and Das Gupta, 1959a: 626. India. insignis Lutz, 1913: 51. Brazil. inamollae Fox and Hoffman, 1944: 110. Puerto Rico (USA). painteri Fox, 1946a: 257. Honduras. isoregalis Majumdar and Das Gupta, in Majumdar et al., 1997: 28. India. jimmiensis Tokunaga, 1959: 229. Papua New Guinea. kinabaluensis Wirth and Hubert, 1989: 211. Indonesia. klossi Edwards, 1933c: 252. Malaysia. lansangensis Howarth, 1985: 58. Laos. liui Wirth and Hubert, 1961: 20. Taiwan. lutzi Costa Lima, 1937a: 419. Brazil. malayae Macfie, 1937d: 471. Malaysia. maruim Lutz, 1913: 48. Brazil. recifei Barbosa, 1947: 25. Brazil. neoregalis Majumdar and Das Gupta, in Majumdar et al., 1997: 29. India. novaguineanus Tokunaga, 1959: 223. Indonesia. ocumarensis Ortiz, 1950b: 455. Venezuela. orestes Wirth and Hubert, 1989: 222. Malaysia. palpalis Macfie, 1948: 78. Mexico. parabubalus Wirth and Hubert, 1989: 224. Malaysia. paraignacioi Spinelli, in Spinelli et al., 1993: 66. Colombia. paraliui Das Gupta, 1962a: 538. India. paramalayae Wirth and Hubert, 1989: 226. Malaysia. paramaruim Wirth and Blanton, 1973: 443. Brazil. pararegalis Majumdar and Das Gupta, in Majumdar et al., 1997: 28. India. parauapebensis Trindade and Felippe-Bauer, 2011b: 42. Brazil. peregrinus Kieffer, 1910: 191. India. judicandus Bezzi, 1916: 8 (1917: 108). Philippines. esmoneti Salm, 1917b: 136. Indonesia. philippinensis Kieffer, 1921g: 564. Philippines. assamensis Smith and Swaminath, 1932: 183 (as variety of peregrinus). India. quadratus Tokunaga, 1951: 108. Indonesia. pikongkoi Howarth, 1985: 51. Laos. plaumanni Spinelli, in Spinelli et al., 1993: 69. Argentina. polypori Wirth and Blanton, 1968a: 212. Panama. pseudodiabolicus Fox, 1946a: 256. Trinidad and Tobago. pseudoheliconiae Felippe-Bauer, in Felippe-Bauer et al., 2008a: 260. Peru.

18 pseudoregalis Majumdar and Das Gupta, in Majumdar et al., 1997: 30. India. quasiregalis Majumdar and Das Gupta, in Majumdar et al., 1997: 30. India. recurvus Delfinado, 1961: 663. Philippines. regalis Majumdar and Das Gupta, in Majumdar et al., 1997: 31. India. ruizi Forattini, 1954a: 189. Brazil. spiculae Howarth, 1985: 53. Laos. subregalis Majumdar and Das Gupta, in Majumdar et al., 1997: 31. India. sumatrae Macfie, 1934c: 190 (1934d: 215). Malaysia. amamiensis Tokunaga, 1937a: 325. Japan. kagiensis Tokunaga, 1937a: 327. Taiwan. ohmorii Takahashi, 1958: 113 (as subspecies of amamiensis). Japan. assimilis Delfinado, 1961: 660. Philippines. tenuifasciatus Wirth and Hubert, 1989: 243. Malaysia. tidwelli Spinelli, in Spinelli et al., 1993: 74. Colombia. travassosi Forattini, 1957: 198. Brazil. trimaculipennis Wirth and Hubert, 1989: 245. Malaysia. trinidadensis Hoffman, 1925: 286. Trinidad and Tobago. oliveri Fox and Hoffman, 1944: 108. Haiti. wokei Barbosa, 1947: 28 (preoccupied by Culicoides wokei Fox, 1947). Panama. diminutus Barbosa, 1951: 163. New name for wokei. venustus Hoffman, 1925: 290. USA (Maryland). verecundus Macfie, 1948: 76. Mexico. xanifer Wirth and Blanton, 1968a: 210. Panama. Subgenus JILINOCOIDES Chu JILINOCOIDES Chu, 1983: 28 (as subgenus of Culicoides). Type species: Culicoides dunhuaensis Chu, by original designation. dunhuaensis Chu, 1983: 28. China. guangxiensis Liu and Hao, 2003: 139. China. mihunensis Chu, 1983: 29. China. minimaporus Liu and Yu, in Yu et al., 2006: 890. China. Subgenus MACFIELLA Fox MACFIELLA Fox, 1955: 217 (as subgenus of Culicoides). Type species: Ceratopogon phlebotomus Williston, by original designation. phlebotomus (Williston), 1896: 281 (Ceratopogon). St. Vincent. amazonius Macfie, 1935a: 52. Brazil. willistoni Wirth and Blanton, 1953c: 116. Panama. Subgenus MARKSOMYIA Bellis and Dyce MARKSOMYIA Bellis and Dyce, 2011: 36 (as subgenus of Culicoides). Type species: Culicoides marksi Lee and Reye, by original designation. dycei Lee and Reye, 1953: 390. Australia (New South Wales). kayi Bellis and Dyce, 2011: 46. Australia (Western Australia). marksi Lee and Reye, 1953: 392. Australia (New South Wales). parvimaculatus Lee and Reye, 1953: 391. Australia (New South Wales). pseudostigmatus Tokunaga, 1959: 234. Indonesia. zentae Bellis and Dyce, 2011: 44. Australia (Queensland).

19 Subgenus MATAEMYIA Vargas MATAEMYIA Vargas, 1960: 43 (as subgenus of Culicoides). Type species: Culicoides mojingaensis Wirth and Blanton, by original designation. albuquerquei Wirth and Blanton, 1973: 424. Brazil. aldomari Felippe-Bauer and Trindade, in Felippe-Bauer et al., 2013:56. Brazil. avilaensis Oritz and Mirsa, 1951: 593. Venezuela. azureus Wirth and Blanton, 1959: 377. Panama. barthi Tavares and Souza, 1978: 619. Brazil. bricenoi Ortiz, 1951a: 445. Venezuela. cuiabai Wirth, 1982b: 250. Brazil. dalessandroi Wirth and Barreto, 1978: 556. Colombia. daviesi Wirth and Blanton, 1968b: 251. Guyana. dicrourus Wirth and Blanton, 1955b: 123. Panama. discrepans Ortiz and Mirsa, 1951: 595. Venezuela. felippebauerae Spinelli, in Spinelli et al., 2007: 660. Brazil. huaynacapaci Felippe-Bauer, in Felippe-Bauer et al., 2008b: 308. Peru. lenti Tavares and Luna Dias, 1980: 396. Brazil. macieli Tavares and Ruiz, 1980: 29. Brazil. mojingaensis Wirth and Blanton, 1953b: 232. Panama. sherlocki Felippe-Bauer and Trindade, in Felippe-Bauer et al., 2013:56. Brazil. volcanensis Wirth and Blanton, 1959: 389. Panama. wallacei Wirth and Blanton, 1973: 449. Brazil. Subgenus MEIJEREHELEA Wirth and Hubert MEIJEREHELEA Wirth and Hubert, 1961: 23 (as subgenus of Culicoides). Type species: Ceratopogon guttifer de Meijere, by original designation. arakawai (Arakawa), 1910: 411 (Ceratopogon, as arakanae). Japan. sugimotonis Shiraki, 1913: 286. Taiwan. alboguttatus Kieffer, 1921g: 563. Taiwan. shimai (Sasaki), 1928: 687 (Ceratopogon). Japan. daleki Smith and Swaminath, 1932: 185. India. micropunctatus Tokunaga, 1951: 105. Indonesia. niigataensis Takahashi, 1958: 115 (as variety of arakawae). Japan. distinctipennis Austen, 1912: 101. Nigeria. multiguttatus (Goetghebuer), 1935d: 156 (Forcipomyia). Democratic Republic of the Congo. guttifer (de Meijere), 1907: 209 (Ceratopogon). Indonesia. hegneri Causey, 1938: 402. Thailand. hildae Cornet and Nevill, 1979: 179. South Africa. histrio Johannsen, 1946: 190 (as variety of guttifer). Guam (USA). mackayensis Lee and Reye, 1953: 383. Australia (Queensland). isechnoensis Glick, 1990: 108. Kenya. leucostictus Kieffer, 1911c: 340. Seychelles. praetermissus Carter, Ingram and Macfie, 1920: 240. Ghana. egypti Macfie, 1924: 66 (as variety of distinctipennis). Egypt. pharao Kieffer, 1925e: 259. Egypt. magnithecalis Majumdar and Das Gupta, in Gangopadhyay and Das Gupta, 2000: 124. India. prolixipalpis Wirth and Hubert, 1989: 426. Malaysia. pycnostictus Ingram and Macfie, 1925: 284. Malawi. alexis de Meillon, 1936: 147. South Africa.